ECOTROPICA 14: 121–128, 2008
© Society for Tropical Ecology
Resuspended intertidal microphytobenthos as
major diet component of planktivorous Atlantic
anchoveta Cetengraulis edentulus (Engraulidae)
from equatorial mangrove creeks
Uwe Krumme1*, Heike Keuthen2, Mario Barletta3, Ulrich Saint-Paul1 & Wolfgang Villwock2
2
1
Center for Tropical Marine Ecology (ZMT), Fahrenheitstr. 6, 28359 Bremen, Germany.
Zoological Institute and Zoological Museum, University of Hamburg, Martin-Luther-King-Platz 3,
20146 Hamburg, Germany.
3 Laboratory of Ecology and Management of Estuarine and Coastal Ecosystems,
Departemento de Oceanografia – UFPE, Cidade Universitaria, Recife, Pernambuco, Brazil
Abstract. In the present study we examined the stomach contents of 49 Atlantic anchoveta, Cetengraulis edentulus (Engraulidae), caught with a block net from intertidal mangrove creeks at diurnal neap tides between June and September 1997
near Bragança (Pará, northern Brazil). On average the fish had eaten 0.53 % ± 0.35 SD of their wet weight, with fuller
stomachs in June/July. Cetengraulis edentulus is a phytoplanktivorous filter-feeder. The principal food was composed of
about half pelagic and half benthic diatoms, predominantly within a 20 to 200 μm size range. Coscinodiscus, other centric
diatoms, and Nitzschia were abundant in the diet throughout the sample period. Sediment particles and detritus were also
ingested. Our observations suggest that foraging occurs close to the substrate and throughout the water column in shallow
turbid waters. The study indicates the importance of intertidal microphytobenthos production during low tide being
made available to transient planktivorous fish at high tide by tidal resuspension. Atlantic anchoveta may occupy a central
position in the food web of many Neotropical mangrove ecosystems along the Atlantic coast. Its close association with
mangroves may partially explain its irregular distribution between Mexico and southern Brazil. Accepted 25 November
2008.
Key words: Bacillariophyta, benthic microalgae, Bragança, food analysis, Pará, phytoplankton, tidal resuspension, tropical mangrove creek.
Introduction
In estuaries, intertidal microphytobenthos resuspensed by tidal currents and waves often makes its way
into food webs through suspension- and depositfeeders (de Jonge & van Beusekom 1992, 1995,
Middelburg et al. 2000, Kang et al. 2006). Phytoplanktivorous fish may also use resuspended microphytobenthos and thus transfer benthic production
to the pelagic food web. However, the role fish play
in linking benthic production to pelagic estuarine
food webs has rarely been studied. Melville & Connolly (2005) found that the food webs supporting
fish on subtropical mudflats were not based on microphytobenthos. The contribution of microphytobenthos exceeded 50% in only one out of 22 species.
If the microphytobenthos production supports fish,
* e-mail: [email protected]
the support is usually indirect via the food web (e.g.,
Sullivan & Moncreiff 1990). However, benthic microalgae are also directly ingested by certain fish
species. Young Mugilidae in particular are known to
feed directly on microphytobenthos from the sediment surface and assimilate it (Lin et al. 2007).
The Atlantic anchoveta Cetengraulis edentulus
(Cuvier, 1829) may provide support for the hypothesis that benthic sources of production are transferred to the pelagic food web by filter-feeding fish.
C. edentulus inhabits inshore, brackish coastal waters
of the western Atlantic between 23°N and 28°S
(Whitehead et al. 1988). Generally C. edentulus is
associated with muddy substrate in shallow, turbid
coastal waters (Simpson 1965, Sergipense & Sazima
1995, Gay et al. 2000), thus is often caught adjacent
to mangroves. The fish seems to favor the innermost
areas of bays and estuaries where phytoplankton
densities are highest (Araújo-Silva et al. 2003; Scho121
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Krumme et al.
ries, pers. comm.). Its distribution is irregular between Mexico and southern Brazil (Simpson 1965),
though the reason for this has not yet been clarified.
C. edentulus has a brush-like filter apparatus
composed of numerous (>100) long and closely
adjoining gillrakers (Bornbusch 1988). The welldeveloped gizzard and grooves lining the inner stomach walls (surface area enhancement) render the genus Cetengraulis unique within the family Engraulidae (Harder 1960, Nelson 1984). Simpson (1965)
studied the early life history of C. edentulus in eastern
Venezuela. Several studies, either from southern Brazil or Colombia, concern the estimation of population parameters (Gallo 1993a,b, Clezar et al. 1994,
Souza-Conceição et al. 2005) or changes in spatiotemporal distribution (Clezar et al. 1993, Sergipense
& Sazima 1995, Gay et al. 2000, Araújo-Silva et al.
2003). The closely related Pacific anchoveta C. mysticetus is distributed along the tropical eastern Pacific
coast from Mexico to Peru; the two species are probably descended from one species that was separated
after the formation of the Isthmus of Panama (LoweMcConnell 1987). Currently C. edentulus is commercially landed only by Venezuela. Annual catches
comprise only 500 t (FAO/FIGIS).
Despite the extended latitudinal distribution of
C. edentulus along the Atlantic coast, studies on the
feeding ecology are restricted to the southern edge of
its distribution in subtropical southern Brazil (Goitein 1984, Clezar 1993, Gay 1995, Sergipense et al.
1999, Gay et al. 2002). Giarrizzo et al. (submitted to
Oecologia) found that C. edentulus from another
mangrove estuary in northern Brazil was highly δ13Cenriched, suggesting a strong dependence on algal
carbon sources. To the best of our knowledge, there
are no studies on the feeding ecology of C. edentulus
from the center of its distribution on the tropical
coast of South America or the Caribbean. Sergipense
et al. (1999) concluded that C. edentulus from the
subtropical Sepetiba Bay, Rio de Janeiro, is a planktivorous filter-feeder. The diet was dominated by
centric and pennate diatoms (relative numerical
frequency: 73 %); oscillatorial cyanophyta were also
abundant (21 %). Dinoflagellates contributed only
3 %. Likewise, the diet of C. edentulus from the
Itaipu lagoon, Rio de Janeiro, was dominated by
diatoms (Gay et al. 2002). A high degree of diatoms
of benthic origin is a typical feature of the stomach
contents of filter-feeding Cetengraulis (Bayliff 1963,
Sergipense et al. 1999). However, it is still unclear
whether benthic diatoms are ingested directly from
the substrate or filtered out of the water. Bayliff
(1963) regarded adult C. mysticetus as iliophagous in
view of the similar relative importance of organisms
in bottom samples and in their stomachs. Sergipense
et al. (1999) had no comparative samples and assumed that the food search of C. edentulus occurs close
to the bottom. Gay et al. (2002) found unselective
feeding in C. edentulus and suggested that the fish
exploit the water layers near the sediment. Thus far,
there has been no in situ observation of the feeding
of the genus Cetengraulis, likely due to the high
turbidity of their foraging grounds (Bayliff 1963).
A better understanding of the feeding ecology of
C. edentulus may help to explain the occurrence of
mass mortality events of this species due to harmful
algal blooms (Mijares et al. 1985; Krumme, unpubl.
data).
The aim of the present study is two-fold: (1) to
contribute to broadening our knowledge of fish
feeding ecology, and in particular to a better understanding of the trophic role of C. edentulus, by analyzing the diet of specimens from an equatorial
mangrove estuary, and (2) to provide observations on
foraging behavior in a dynamic tidal environment.
Material & Methods
Study area and study site. The study area is located in
the Caeté River estuary near Bragança, about 200 km
south-east of the Amazon delta. It is part of the
­second largest mangrove area in the world, covering
about 7000 km2 of the north-east coast of South
America (Kjerfve & Lacerda 1993). Annual average
rainfall is about 2500 mm. There is a wet season
(January to June) and a dry season (August to
­November), when estuarine salinities can fall below
5 psu and exceed 35 psu respectively. Water tem­
perature is high year-round, ranging from 27ºC to
30ºC.
The sample sites were three intertidal 1st-order
mangrove creeks in the muddy upper reaches of the
Furo do Meio, located in the central part of the
peninsula bordering the western side of the Caeté Bay
(see Figure 1 in Barletta-Bergan et al. 2002). In the
Caeté estuary, intertidal 1st-order creeks (c. 25 m
wide at the mouth, 3–5 m deep at high water [HW])
that are further ramified upstream, drain the mangrove area into large subtidal main channels (furos)
that connect to Caeté Bay. According to BarlettaBergan (1999), creeks 1, 2, and 3 had 6481, 9351,
and 1865 m2 of inundated area at neap tide HW
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Feeding ecology of Cetengraulis edentulus
respectively. Creek 1 was located upstream of a wooden
bridge crossing the Furo do Meio in its upper reaches,
while creeks 2 and 3 were located downstream of it.
The bridge is c. 50 m wide and 6–7 m high, with
1.5–2 m between poles. The tidal range of the semidiurnal tide is 2–3 m at neap and 3–5 m at spring
tides. The tide is asymmetric; flood and ebb tide last
4 and 8 hours respectively. In the last 4 hours, ebb
tide is extremely weak with an almost negligible fall
in the water level.
Sampling method. C. edentulus were caught for stom­
ach analysis during diurnal neap ebb tides of the last
quarter moon. Samples were taken monthly between
June and September 1997 as part of the one-year
study of Barletta-Bergan et al. (2002) and Barletta et
al. (2003). The three creeks were sampled on consecutive days. Each day at slack HW, a creek mouth
was blocked with a net (50 m × 5 m, 1 cm stretched
mesh size). Fish were collected during ebb tide until
the creek was totally drained (c. six hours after HW).
In the field, specimens were directly fixed in 15 %
borax-buffered formalin in sea water. Salinity increased from 22 to 24.7, 28.7, and 31.5 psu between
June and September respectively.
Length relationships. Total length (TL), standard length
(SL) (both to the lowest 0.5 cm), total wet weight,
and wet weight without stomach were taken (Sartorius LC 4200 S ± 0.01 g). The fish were assigned to
three size classes: < 9 cm, 9- < 11 cm, ≥ 11 cm SL.
The length of the uncoiled digestive tract (DT) was
measured from the pylorus to the anus (± 0.5 cm).
Relative length of the DT (RLDT) was calculated
(DT length × TL-1).
Stomach fullness. The stomach fullness index (SFI)
was calculated as SFI = [(stomach content weight ×
100) × total fish weight-1]. The intestines were transferred to 8 % buffered formalin, washed, drained on
an absorbent paper and wet-weighed (± 0.01 g). The
weight of the contents of the intestines (C) was
­calculated as C = R – E; where R is the weight of the
removed intes-tines (g), E is the weight of the empty
intestine (g).
A two-way analysis of variance (ANOVA) was
calculated to ascertain whether SFI differed by
month and size class (Sokal & Rohlf 1995). To fulfill
the ANOVA assumptions, SFI raw data (y) were
transformed using a logistic transformation [log(y/
(100-y))] which transforms SFI values from the
0–100 scale to the range from plus or minus infinity.
To avoid an incomplete design, the months June/
July and August/September were pooled. The three
creeks were not considered as a source of variance.
The Scheffé test was used for post hoc analysis.
Stomach content analysis. Stomach contents were diluted in distilled water and stirred to make a homogeneous solution. A defined amount of the solution
was transferred with a pi-pette to a microscope slide
for identification of food items to the lowest pos­sible
taxonomic level (Leitz: Dialux microscope) (Hustedt
1965, Newell & Newell 1970, Drebes 1974, Round
1975, Round et al. 1996).
The size categorization of plankton followed Sieburth et al. (1978). The presence of food items was
recorded as abundant, regular, rare, or sporadic. The
origin of the diatom genera ingested was assigned to
either benthic or pelagic according to the literature.
Genera were omitted that occurred only sporadically
or had a vague classification such as Paralia sp. and
Frickea sp. (Round et al. 1996). Each abundance
category was substituted by a value (abundant: 3,
regular: 2, rare: 1) and the monthly percentage of
benthic and pelagic diatoms was determined.
After identification, the stomach contents were
dried (24 hrs at 60°C) and weighed (dry weight =
WD; Sauter Typ 404; ± 0.00001 g). The samples
were burned (24 hrs at 510°C in a muffle furnace)
and weighed (ash free dry weight = WAFD). The difference between WD and WAFD gave the amount of
organic matter (g) that should have been available
for the fish and an unknown proportion of organic
matter already digested.
Results
We caught 9, 8, 15, and 17 C. edentulus in June,
July, August, and September 1997 respectively (n =
49; size range: 7.5–12 cm SL). All but three C. edentulus were caught in the two creeks downstream of
the bridge (creek 2: n = 33, creek 3: n = 13) suggesting that obstacles such as bridge piers standing
closely side by side may restrict the daytime passage
of schools of C. edentulus.
Stomach fullness. The stomach of C. edentulus is extremely furled. The mean RLDT was 4.43 ± 0.85 SD
(range: 2.63–5.37; n = 17). Only four out of 49
specimens (8 %) had an empty stomach. The others
seemed to be well filled. Different stomach fullness
stages could not be determined due to the anatomy
123
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Krumme et al.
of the stomachs and the consistency of the diet. SFI
was significantly higher in June/July (late wet season)
than in August/September (early dry season) while
size class had no significant effect on SFI (two-way
ANOVA, month: df: 1, p < 0.05; size class: df: 2, p
> 0.30; interaction: df: 2, p > 0.08). On average the
fish had eaten 0.53 % ± 0.35 SD of their wet weight
(n = 49; range: 0.07–1.55 %).
Diet composition. The consistency of the stomach
contents of C. edentulus resembled a clayey compressed matter of low water content. Twenty-one
genera were identified. The diet consisted almost
exclusively of phytoplankton, mainly diatoms (Table
1). Coscinodiscus sp., other centric diatoms and Nitzschia sp. were abundant throughout the sample period. Sceletonema costatum and Navicula sp. were
only abundant in June. In contrast, Raphoneis suri­rella
(abundant), Thalassionema sp. (regular), Odontella sp.
(rare), and Paralia sp. (rare) occurred mainly from
July to September. Dinoflagellates were rare in June/
July and sporadic in August/September. The stom­
achs also contained detritus and sediment particles.
Size composition. The size of most plankton orga­
nisms was between 20 μm and 200 μm, i.e., micro-
Table 1. Abundance and sizes of food items of C. edentulus between June and September 1997 from mangrove creeks near Bragança, Pará, northern Brazil. C: Centrales, Pe: Pennales. Origin: Assignment of taxonomic groups to B: Benthos, P: Plankton. ++++: abundant, +++: regular, ++: rare, +: sporadic, ---: absent.
Taxa, genera or type
Order Origin
Abundance
June July
BACILLARIOPHYCEAE
Coscinodiscus sp.
C
+++
remaining Centrales
Sceletonema costatum
C
C
Odontella sp.
C
Paralia sp.
Ditylum cf. brightwellii
Chaetoceros sp.
Navicula sp.
Raphoneis aff. surirella
Nitzschia sp.
C
C
C
Pe
Pe
Pe
Thalassionema sp.
Pe
Pleurosigma sp. and Gyrosigma sp.
Pe
Frickea sp.
Bacillaria sp.
Diploneis sp.
Pe
Pe
Pe
Delphineis sp.
Cymatosiraceae cf. Campylosira sp.
Pe
Pe
cf. Stenopterobia sp.
Pe
Haslea sp.
Pe
spec. E (cf. Eunotogramma sp.)
(C)
spec. A
Pe
CYANOBACTERIA cf. Oscillatoria sp.
DINOFLAGELLATA
SILICOFLAGELLATA
OTHERS: Veliger larva
Nematoda
P
Diameter, or length × width (µm)
Aug. Sept.
+++
++++ +++261; 221; 208; 186; 166; 102; 97;
69; 50; 14
P/B +++ ++++ ++++ +++ 69; 66; 52; 44; 38; 33; 30; 25; 16
P
++++ ++
++
+++Chain: 22×13; 20×12; 16×11; 14×10;
19×8
P
+
++
++
++200×91; 111×100; 44×19; 36×25;
33×22; 27×16
P/B (?) ---
++
++
++
Chain : 13×33; 20×36; 7×10; 7×11
P
++
+++ ++
+
P
---
---
---
+
Chain: 8×3
B
++++ +++ ++
+++ 47×8; 36; 27; 17; 16; 14; 11
B
++
++++ ++++ ++++ Chain: 27×4; 16×3; 14×3; 11×3; 8×4
P
++++ +++ ++++ ++++220; 158×10; 155; 147; 130; 119;
97; 40; 36; 16
B
++
+++ +++ +++172×6; 169×6; 136×6; 122×6; 113×6;
97×6; 75×4; 44×3
B
++
++
++
++155×14; 91×8; 75×8; 58×8; 50×10;
36×4
?
+
---
+
+
100
B
---
---
+
+
Colony: 122×4; 77×4; 72×4
B
---
++
++
++60×22; 52×20; 44×19; 36×13; 33×13;
30×11
B
---
+
+
+
30×3; 20×3
(B)
---
---
++
++Chain: 36×6; 25×5; 24×4; 22×4;
20×4
B
---
---
---
+
341; 332; 266; 122
B
++
++
++
+
127×6; 69×4; 38×2
(P)
---
---
+
+
47×52; 44×55; 44×52
?
+
+
+
++
83×41; 58×27
P
---
+
+
+
Chain: 22×33; 3×11
P
++
++
+
+
41×19; 41×22; 36×22
P
---
---
+
+
27; 4
P
---
---
+
---
27×36
B
---
+
---
---
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Feeding ecology of Cetengraulis edentulus
phytoplankton (range: 4–341 μm) (Table 1). Chains
usually consisted of only three to six cells. Only
Navicula sp. and silico-flagellates were assigned to the
nanoplankton (< 20 μm). Larger than 100 μm were
Coscinodiscus sp., Odontella sp., Thalassionema sp.,
Bacillaria sp., Gyrosigma sp., Nitzschia sp., Frickea
sp., Haslea sp. and Stenopterobia sp. Only Stenopterobia sp. and Coscinodiscus sp. were > 200 μm and
­assigned to the mesoplankton (0.2–20 mm).
Benthic vs. pelagic diatoms. Out of a total of 21 gen­era
identified, 12 were assigned to a benthic and nine to
a pelagic origin. Diatoms from both the benthic and
the pelagic compartment accounted for approxi­
mately 50 % of the diet throughout the sample
­period.
Organic content. The mean organic matter content of
the stomach contents was 44.5 % ± 10.1 SD (range:
28.6–73.8 %, n = 44). The organic matter content
did not differ between the four months (KruskalWallis test, n = 44, p > 0.32).
Discussion
The Atlantic anchoveta C. edentulus is a planktivorous
primary consumer, filter-feeding on primarily centric
and pennate diatoms. A comparison of the diet of C.
edentulus from the tropics (present study) and the
subtropics (Sergipense et al. 1999) shows that the
fractions of the principal phytoplankton groups in
the diet were similar. However the taxonomic composition and the fraction of different species within
the phytoplankton groups vary. Such differences in
the list of food items may be related to sample size,
sampling season, and preciseness of identification.
The diet of C. mysticetus in the eastern Pacific was
also dominated by centric and pennate diatoms, while
other organisms were of insignificant importance
(Bayliff 1963). The similarities in the diets of C.
edentulus and C. mysticetus suggest that these sister
species have similar foraging strategies and occupy
similar ecological niches.
Tidal currents disperse benthic diatom mats. It is remarkable that benthic diatoms were just as abundant
in the diet of C. edentulus as pelagic diatoms. A
comparison of the diet composition of C. edentulus
with the results from surface phytoplankton surveys
by Schories (unpubl. data) and Dummermuth (1997)
suggests that the relative contribution of benthic and
pelagic diatoms was similar in the water column and
in the stomachs. Schories (unpubl. data) studied the
tidal and diel dynamics of the phytoplankton community in the Furo do Meio during twelve months
in 1996/1997, thus overlapping in time with the
present study. Dummermuth (1997) investigated the
primary production of the phytoplankton and the
microphytobenthos during three dry season months
in 1996 in the same area. The species abundant in
the two studies (S. costatum, Nitzschia sp., Navicula
sp., and R. surirella) were also abundant in the stom­
achs of C. edentulus. Despite its low abundance in
the Furo do Meio, Coscinodiscus was categorized as
abundant in the stomachs. This asymmetry could be
due to a positive selection by C. edentulus or to an
abundance overestimation of the large Coscinodiscus
cells during the stomach content analysis.
Benthic diatoms play a significant role in the
pelagic food web of the furos (Schories, pers. comm.)
because the strong tidal currents disperse the upper
sediment layers at each tide (Lucas et al. 2000), with
current speeds < 0.5 m s-1 at neaps and often > 1.2 m
s-1 at springs in the study area. The ability of the tidal
current to displace surface sediments even at weak
neap tides was observed in the Furo do Meio, when
ADCP measurements were unusable because bedload
transport of fine muddy substrate led to missing
bottom echoes (Krumme & Hanning 2005). The
average Secchi depth is only 0.3 m (Krumme, unpubl. data) and seston content is high, commonly
around 0.1 g l-1 (Schories, unpubl. data).
Pickney & Zingmark (1991) showed that intertidal benthic diatoms undergo regular vertical migrations to the surface for photosynthesis during the
low-tide period. During the low-tide period in the
furos, extensive mats of benthic diatoms appear on
the sediment surface of flat sections and slip-off
slopes of exposed intertidal muddy banks, especially
in the upper reaches of the furos (Dummermuth
1997). Although the majority of the benthic diatoms
may slide down sufficiently deep into the sediment
prior to inundation at flood tide, the rapid first flood
rise (Krumme 2004) regularly flushes away remains
of the mats (Dummermuth 1997), together with fine
sediment particles. This resuspension of benthic diatoms from the muddy intertidal increases the overall
abundance of “pelagic” phytoplankton, making the
furos the areas with the highest phytoplankton abundance in the Caeté estuary (Schories, pers. comm.).
The exceptional nature of this phenomenon in the
upper reaches may be the reason for C. edentulus
foraging in this section of the furos.
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Krumme et al.
Foraging habits and sediment particles in the diet.
Most C. edentulus stomachs from the intertidal creeks
contained food and were well filled, suggesting that
C. edentulus entered the creeks to feed during the
daytime HW period. Sergipense et al. (1999) also
found filled stomachs in daytime samples. However,
C. edentulus likely feeds continuously. The stomach
fullness of C. edentulus sampled during consecutive
neap tide cycles in the Furo do Meio suggested
continuous feeding throughout the tidal cycle, both
day and night (Krumme 2004). Algivores/detrivores
rarely have empty stomachs because they have relatively short feeding intervals on account of the low
energy content, small particle size, and low conversion efficiency of their food (Brett & Groves 1979,
Arrington et al. 2002).
The great abundance of benthic diatoms in stomachs of C. edentulus caught with beach seines in
shallow water led Sergipense et al. (1999) and Gay
et al. (2002) to assume that C. edentulus filter-feed
close to the bottom. However, C. edentulus probably
feed throughout the water column because strong
tides disperse benthic diatoms. During the hydroacoustic observation of fish movements in the Furo
do Meio (Krumme & Saint-Paul 2003), fish schools
were seen beneath the water surface (0–2 m), suggesting that C. edentulus forage throughout the water
column in the relatively shallow furos (< 5 m deep
during low tide). In fact, C. edentulus seem to feed
anytime and everywhere. They were seen filtering
with mouths wide open day and night, at flood and
ebb tide, and in creeks and channels at water depths
ranging from 0.5 m to 5 m (Krumme, pers. obs.).
Bayliff (1963) mentioned that C. mysticetus may
also stir up the muddy sediment while foraging,
probably a behavior used when “natural” phytoplankton concentrations are too low due to lack of organism resuspension by tides or waves. Such an active
disturbance of the uppermost sediment may be used
by C. edentulus at the banks of shallow channels
during the stagnant low-tide phase when the phy­
toplankton falls. During low tide, Krumme et al.
(2004) caught C. edentulus in the Furo in shallow
water < 1.5 m deep, where artisanal fishermen often
stumble across agglomerations of C. edentulus while
hand-trawling for penaeid prawns.
The sediment particles in C. edentulus are almost
inevitably ingested while filter-feeding in turbid waters. The organic matter content of the sediment
particles in C. edentulus stomachs (44.5 %) was much
higher than (e.g.) in iliophagous Mugil cephalus (6–
10 %) (Odum 1970). However, though not quantified, the relative contribution of mineral material to
the total stomach content was certainly much lower
than in M. cephalus (40–100 %). The dominance of
diatoms in the diet, the foraging habit, and the low
proportion of sediment particles in the stomachs
render it unlikely that C. edentulus is as iliophagous
as Bayliff (1963) concluded for adult C. mysticetus.
The sediment particles in the stomach likely function
as an agent to grind diatoms, and may facilitate the
function of the gizzard of C. edentulus similar to M.
cephalus (Odum 1970).
Conclusion. The Antlantic anchoveta is a plankti­
vorous filterfeeder, predominantly of diatoms. The
study provides evidence for the importance of intertidal microphytobenthos production during low tide
being made available to a transient phytoplankti­
vorous fish at high tide by tidal resuspension. The
diet of C. edentulus seems to be both unaffected by
latitude and similar to the sister species C. mysticetus.
The foraging activity of C. edentulus seems to be
restricted to shallow muddy areas close to mangroves
(e.g., Clezar et al. 1993, Sergipense & Sazima 1995,
Araújo-Silva et al. 2003) where significant amounts
of benthic diatoms are added to the pelagic phytoplankton, either passively by tidal currents and wave
action or actively by the fish stirring up the muddy
substrate themselves. Therefore the presence of mangroves and occurrence of C. edentulus may be closely
correlated and may partially explain its irregular
distribution between Mexico and southern Brazil
(Simpson 1965). This would further highlight the
central trophic position the primary consumer C.
edentulus may have in the food webs of many Neotropical mangrove ecosystems on the Atlantic coast.
Given its important trophic role, more information
on the spatio-temporal dynamics of foraging, distribution, and autotrophic sources of C. edentulus is
urgently required.
Acknowledgments
We are grateful to Audrey Barletta-Bergan and the
fishermen Walmiro and André for support during the
sampling. This work resulted from the cooperation
between the Center for Tropical Marine Ecology
(ZMT), Bremen, Germany, and the Univ. Federal do
Pará (UFPa), Belém, Brazil, under the Governmental
Agreement on Cooperation in the Field of Scientific
Research and Technological Development between
Germany and Brazil financed by the German Mi-
126
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Feeding ecology of Cetengraulis edentulus
nistry for Education, Science, Research and Technology (BMBF) [Project number: 03F0253A, Mangrove Dynamics and Management – MADAM], and
the Conselho Nacional de Pesquisa e Tecnologia
(CNPq) [MADAM contribution 119].
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